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An Examination of Feeding Ecology in Pleistocene Proboscideans from Southern China (Sinomastodon, Stegodon, Elephas), by Means of Dental Microwear Texture Analysis

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An Examination of Feeding Ecology in Pleistocene Proboscideans from Southern China (Sinomastodon, Stegodon, Elephas), by Means of Dental Microwear Texture Analysis Quaternary International xxx (2016) 1e11 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint An examination of feeding ecology in Pleistocene proboscideans from southern China (Sinomastodon, Stegodon, Elephas), by means of dental microwear texture analysis * Hanwen Zhang a, Yuan Wang b, c, , Christine M. Janis a, Robert H. Goodall d, ** Mark A. Purnell d, a School of Earth Sciences, University of Bristol, Wills Memorial Building, Queen's Road, Bristol BS8 1RJ, UK b Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China c State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China d University of Leicester, Department of Geology, Leicester LE1 7RH, UK article info abstract Article history: It has long been thought that environmental perturbations were the key driving force behind the suc- Available online xxx cession of three distinct mammal faunas in the Pleistocene Ailuropoda-Stegodon faunal complex (sensu lato) of South China: the lower Pleistocene Gigantopithecus-Sinomastodon fauna, the middle Pleistocene Keywords: Ailuropoda-Stegodon fauna (sensu stricto) and the upper Pleistocene Homo-Elephas fauna. Here, we apply Pleistocene proboscideans three-dimensional dental microwear texture analysis (DMTA) to three characteristic fossil proboscideans South China from these mammal faunas to provide preliminary tests for hypotheses of trophic ecology. Despite a few DMTA methodological caveats, this study demonstrates the potential of DMTA for understanding the diets of Microwear fl Feeding ecology fossil proboscideans. The texture of microwear in Sinomastodon and Stegodon are more re ective of browsing, whereas that of Elephas is suggestive of mixed feeding. The results suggest a more complex process of Pleistocene faunal turnovers in South China. Rather than a unidirectional trend of climate- driven environmental deterioration, biotic factors might have played a more substantial role than pre- viously thought. © 2016 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). 1. Introduction order (Shoshani and Tassy, 1996, 2005; Sukumar, 2003). With a comprehensive proboscidean phylogeny still a formidable task It has been widely recognised that the evolution and turnover of (Shoshani, 1998; Adrian Lister, personal communication), it is megaherbivores during the Plio-Pleistocene was fundamentally crucial to understand their palaeobiology with respect to the governed by climate changes on local and global scales (Qiu, 2006; environment, as the proboscideans are often deemed to be typical Wei et al., 2010; Wang et al., 2012; Fortelius et al., 2014). This is K-selected large mammal species, with sensitive responses to cli- particularly evident in the Proboscidea, whose fossil record in- matic and environmental fluctuations. dicates a major decline in diversity throughout this period, even- It mayappear superficially that the proboscideans conform to the tually leaving only three extant species of a previously very diverse classic narrative of large herbivore evolution during the Cenozoic, of turning from browsing to grazing. As the global climate gradually deteriorated, forests and woodlands transformed into more arid grasslands towards the late Cenozoic (Janis, 1993, 2008), and many * Corresponding author. Institute of Vertebrate Paleontology and Paleoanthro- large herbivores evolved dentitions capable of processing more pology, Chinese Academy of Sciences, Xizhimen-Wai Street 142, Beijing 100044, China. abrasive vegetation (see Damuth and Janis, 2011 for detailed re- ** Corresponding author. University of Leicester, Department of Geology, Leicester view). The proboscideans saw the transition from the bunolopho- LE1 7RH, UK. dont morphology of the molars in the broadly ancestral E-mail addresses: [email protected] (Y. Wang), [email protected] “gomphotheres”, to lophodonty in two families, Stegodontidae and (M.A. Purnell). http://dx.doi.org/10.1016/j.quaint.2016.07.011 1040-6182/© 2016 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). Please cite this article in press as: Zhang, H., et al., An examination of feeding ecology in Pleistocene proboscideans from southern China (Sinomastodon, Stegodon, Elephas), by means of dental microwear texture analysis, Quaternary International (2016), http://dx.doi.org/ 10.1016/j.quaint.2016.07.011 2 H. Zhang et al. / Quaternary International xxx (2016) 1e11 Elephantidae, as shown in Fig. 1 (although the precise in- super-keystone species in their ecosystem (Fritz et al., 2002), but terrelationships remain weakly resolved: see Sanders et al., 2010, also because studies have shown that both fossil and extant pro- and references therein). Stegodonts retained the brachyodonty of boscideans are highly generalist herbivores whose dietary gomphotheres, but the cusps and lophs which typify the bunolo- composition reflects local environmental conditions more than phodont morphology were multiplied to form enamel ridges more their “adaptive optimal” diet (e.g. Rivals et al., 2012, 2015; Lister, optimised for shearing (Fig. 1), similar to those seen in extant ele- 2013, see discussion below). China has an enormous abundance phants (Saegusa, 1996; von Koenigswald, 2014). Evidence from and diversity of large mammal fossils from the late Cenozoic, morphology, geochronology and palaeobiogeography indicate that including proboscideans (e.g. Owen, 1870; Hopwood, 1935; Colbert elephantids like Elephas evolved lophodonty in parallel with steg- and Hooijer, 1953; Chow and Zhang, 1974; Han, 1987; Pei, 1987; Qiu odonts such as Stegodon (Saegusa et al., 2005, 2014; Wang et al., et al., 2004; Qiu, 2006; Wang et al., 2013; Chi and Wei, 2014; Jin 2015a; Ao et al., 2016). Elephantids have fused the transverse cusp et al., 2014; Zeitoun et al., in press): a long history of research of pairs of the gomphothere molars into continuous loops of enamel, these materials has also provided an established literature for un- each filled with dentine and joined to its neighbours by cement derstanding the process of proboscidean evolution and turnover in (Rivals et al., 2015). In more derived elephantids, the tooth crown is the context of environmental change and faunal turnover (Maglio, also dramatically heightened (Maglio, 1973; also see Fig. 1). 1972; Pei, 1987; Chen, 1999, 2011; Tong, 2007; Jin et al., 2009a,b; The bunolophodontelophodont transition in the proboscidean 2014; Wei et al., 2010; Wang et al., 2012; Button et al., 2013, 2014b). fossil record (Fig. 1), and the independent evolution of hypsodonty Recently, chronologically well-constrained Pleistocene mammal (increased height of tooth crown) among the different elephantid fossil-bearing strata, have been reported from karst caves and fis- genera (Maglio, 1973), has long been linked with a transition from sures of the Guangxi Zhuang Autonomous Region in southernmost browsing to grazing (Savage and Long, 1986; Shoshani, 1998). China. Systematic survey of biostratigraphy across areas in the vi- However, recent application of more direct dietary proxies allow cinity of Chongzuo Ecological Park and the Bubing Basin has palaeontologists to decouple phenetic and behavioural inferences, enabled the definition and refined timescale of three key faunal in order to better interpret the evidence for evolutionarily adaptive stages making up South China's characteristic Ailuropoda-Stegodon responses in the fossil record (Lister, 2014). Cerling et al. (1999) faunal complex (sensu lato, but see Zeitoun et al., in press) during demonstrated, using stable carbon isotope signatures of the the Pleistocene: the early Pleistocene Gigantopithecus-Sinomasto- dental enamel, that despite the C4 signal of a grazing-dominated don fauna, the middle Pleistocene Ailuropoda-Stegodon fauna (sensu diet for the last 7 Ma, African proboscideans in the late Pleisto- stricto) and the late Pleistocene Asian elephant fauna, corroborated cene showed a reversal to a signal of C3 mixed-feeding, this is in by an abundance of other Pleistocene fossils from Guangxi (Jin et al., spite of a continuing trend towards hypso-lophodonty (Maglio, 2007, 2009a; 2014; Wang et al., 2007, 2014a,b; Rink et al., 2008). 1973; Lister, 2013). Detailed corroboration of isotopic proxies for The fossil proboscideans (Sinomastodon, Stegodon, Elephas)were grazing and grassland domination with the African proboscidean always typical members of Quaternary faunas of southern China fossil record suggests a time lag between emergence of grazing (Zeitoun et al., 2015, in press). behaviour and subsequent morphological response in the appear- Sinomastodon, the terminal and only brevirostrine Old World ance of a hypso-lophodont dentition (Lister, 2013). In fact, the trilophodont gomphothere, possesses typical brachyo- gomphotheres remained the dominant grazing proboscideans for bunolophodont molars (Fig. 1)(Tobien et al., 1986; Chen, 1999; much of the late Cenozoic in Africa (Cerling et al.,1999; Lister, 2013). Wang et al., 2012, 2014b). The fossil record of Sinomastodon The focus of this study is the proboscidean succession in the shows a northward shift during the latest
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